View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by ESC Publications - Cambridge Univesity Biol. Rev. (2014), pp. 000–000. 1 doi: 10.1111/brv.12168 Making sense of ‘lower’ and ‘upper’ stem-group Euarthropoda, with comments on the strict use of the name Arthropoda von Siebold, 1848 Javier Ortega-Hernández∗ Department of Earth Sciences, University of Cambridge, Downing Site, Cambridge, CB2 3EQ, U.K. ABSTRACT The ever-increasing number of studies that address the origin and evolution of Euarthropoda – whose extant representatives include chelicerates, myriapods, crustaceans and hexapods – are gradually reaching a consensus with regard to the overall phylogenetic relationships of some of the earliest representatives of this phylum. The stem-lineage of Euarthropoda includes numerous forms that reflect the major morphological transition from a lobopodian-type to a completely arthrodized body organization. Several methods of classification that aim to reflect such a complex evolutionary history have been proposed as aconsequence of this taxonomic diversity. Unfortunately, this has also led to a saturation of nomenclatural schemes, often in conflict with each other, some of which are incompatible with cladistic-based methodologies. Here,I review the convoluted terminology associated with the classification of stem-group Euarthropoda, and propose a synapomorphy-based distinction that allows ‘lower stem-Euarthropoda’ (e.g. lobopodians, radiodontans) to be separated from ‘upper stem-Euarthropoda’ (e.g. fuxianhuiids, Cambrian bivalved forms) in terms of the structural organization of the head region and other aspects of overall body architecture. The step-wise acquisition of morphological features associated with the origins of the crown-group indicate that the node defining upper stem-Euarthropoda is phylogenetically stable, and supported by numerous synapomorphic characters; these include the presence of a deutocerebral first appendage pair, multisegmented head region with one or more pairs of post-ocular differentiated limbs, complete body arthrodization, posterior-facing mouth associated with the hypostome/labrum complex, and post-oral biramous arthropodized appendages. The name ‘Deuteropoda’ nov. is proposed for the scion (monophyletic group including the crown-group and an extension of the stem-group) that comprises upper stem-Euarthropoda and Euarthropoda. A brief account of common terminological inaccuracies in recent palaeontological studies evinces the utility of Deuteropoda nov. as a reference point for discussing aspects of early euarthropod phylogeny. Key words: Euarthropoda, Panarthropoda, Tardigrada, Onychophora, Deuteropoda nov., phylogeny, crown-group, lobopodian, arthrodization, Aiolopoda. CONTENTS I. Introduction .................................................................................................... 2 II. Arthropoda, Euarthropoda, or Panarthropoda? A note on nomenclature ............................. 2 (1) The status of ‘Lobopodia Snodgrass, 1938’ ............................................................ 5 (2) Suggested taxonomic consensus ......................................................................... 7 III. Diversity in stem-group Euarthropoda ...................................................................... 7 IV. Transition from ‘lower’ to ‘upper’ stem-Euarthropoda ................................................... 10 (1) The problem with Schinderhannes ........................................................................ 10 (2) Insights from head segmentation ....................................................................... 12 (a) Considerations for taxa with unusual morphologies ............................................. 13 * Address for correspondence (Tel: +44 (0) 1223 333416; E-mail: [email protected]). Biological Reviews (2014) 000–000 © 2014 Cambridge Philosophical Society 2 Javier Ortega-Hernández V. Comparison with ‘three-phase model of euarthropod evolution’ ....................................... 14 VI. On naming confusion and palaeontological inaccuracies ................................................ 15 VII. A new classification for scion upper stem-Euarthropoda+Euarthropoda .............................. 15 (1) The applicability and fossil record of Deuteropoda nov. ............................................. 16 VIII. Conclusions ..................................................................................................... 17 IX. Acknowledgements ............................................................................................. 17 X. References ....................................................................................................... 18 I. INTRODUCTION and ‘upper’ stem-Euarthropoda based on the results of recent phylogenetic analyses, and propose a clear The application of stem- and crown-group concepts synapomorphy-based definition that can be applied (Jefferies, 1979; see also Budd & Jensen, 2000) has when discussing aspects of early euarthropod evolution. led to an improved understanding of the evolutionary The suggested categorization is then compared to alter- links between extinct and extant organisms, particularly native models that seek to distinguish between different with the widespread usage of parsimony-based cladis- evolutionary stages and taxonomic levels in this phylum. tic methods for exploring phylogenetic relationships. The appropriate usage of systematic and phylogenetic Euarthropoda Lankester, 1904 (see Table 1) – the nomenclature is addressed in light of these discussions monophyletic group including Chelicerata (pycno- in order to rectify recent terminological inaccuracies, gonids, xiphosurans, arachnids) and Mandibulata and also to promote its correct application in future con- (myriapods, crustaceans and hexapods) – represents tributions. a classical example of the utility of this nomenclatural convention. Numerous studies produced in the last two decades indicate that total-group Euarthropoda is II. ARTHROPODA, EUARTHROPODA, OR typified by a long and diverse stem-lineage that includes PANARTHROPODA? A NOTE ON forms characterized by a lobopodian-like construc- NOMENCLATURE tion (i.e. lobopodians, gilled-lobopodians; Fig. 1A, B), partially arthropodized limb sets (i.e. radiodontans; The literature addressing the classification, morphology Fig. 1C), and even fully arthrodized bodies (e.g. fuxi- and evolution of Euarthropoda is rich and often convo- anhuiids, Cambrian bivalved euarthropods; Fig. 1D, E). luted, mainly as a result of its lengthy history and contin- Given that stem-Euarthropoda encompasses disparate uous growth. Given that using precise nomenclature is types of body organization, a further categorization of fundamental for an efficient communication, it is nec- these representatives is desirable for practical reasons, essary to elucidate the origins and applicability of the such as discussing the phylogenetic position of specific more recurrent terminology. As a pre-emptive conven- taxa relative to each other, and describing the sequential tion for clarity, Euarthropoda sensu Lankester (1904) is acquisition of major synapomorphies leading to evolu- used throughout the discussion instead of its senior syn- tionary origins of the crown-group (e.g. Budd, 2002; onym, Condylopodes Bronn 1850 (see Table 1), given Daley et al., 2009; Kühl, Briggs & Rust, 2009; Liu et al., that the former term carries a greater familiarity in the 2011; Legg, Sutton & Edgecombe, 2013; Ma et al., 2014; recent literature (see also Hegna et al., 2013). Smith & Ortega-Hernández, 2014). However, a formal The precise use of the name ‘Arthropoda’ – as well sub-categorization of stem-Euarthropoda is lacking, the taxonomic scope it encompasses – have been the likely due to the scarcity of clearly monophyletic clades source of much nomenclatural confusion (Table 1). within this stem-lineage and the unstable phylogenetic In a recent revision on the term’s authorship, Hegna position of some of its constituent fossil taxa. et al. (2013, p. 72) drew attention to the original diag- Budd (2002, 2008) is so far the only author that nosis of Arthropoda von Siebold, 1848; in addition has suggested a potential categorization by introducing to bilateral symmetry and jointed limbs, Arthropoda the concept of ‘upper stem-group Euarthropoda’; refer- is characterized by the possession of a ring of gan- ences to this term have been used by subsequent work- glia around the oesophagus, followed by a chain-like ers, which suggest the that the utility of such a distinc- (ganglionated) nerve cord. Based on this diagnosis, tion is welcomed (e.g. Budd & Telford, 2009; Daley et al., von Siebold (1848) utilized Arthropoda for a group 2009; Edgecombe, 2009; Legg et al., 2013). However, comprising Crustacea (considered also to include there is no clear definition of upper stem-Euarthropoda Myriapoda), Hexapoda (listed as ‘classis Insecta’) and or its applicability, and thus it has been largely uti- Chelicerata (listed as ‘classis Arachnida’). Significantly, lized as a non-descript term that embodies a super- von Siebold (1848, p. 586) classified Tardigrada (water ficial notion of body organization present in some bears) as a suborder under his ‘classis Arachnida’, Palaeozoic representatives. The aim of this contribu- and assigned the former group the same taxonomic tion is to produce a formal distinction between ‘lower’ rank as Pycnogonida (sea spiders). This context makes Biological Reviews (2014) 000–000 ©